305 Zitteliana 89

Vertebrate fossils from the Kimmeridgian of Brunn: the oldest fauna from the Solnhofen Archipelago (Late Jurassic, Bavaria, Germany)

Paläontologie Bayerische OliverGeo W.Bio- M. Rauhut1,2,3*, Adriana López-Arbarello1,2,3, Martin Röper4,1 & Monika Geobiologie Center & Staatssammlung Rothgaenger4,1 LMU München für Paläontologie und Geologie LMU München

1  München, 01.07.2017 SNSB, Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333 Munich, Germany 2 LMU  Manuscript received GeoBioCenter , Ludwig-Maximilians-University, Richard-Wagner-Str. 10, 80333 Munich, 17.10.2016; revision Germany 3 accepted 05.12.2016 Department of Earth and Environmental Sciences, Ludwig-Maximilians-University, Richard- Wagner-Str. 10, 80333 Munich, Germany  ISSN 0373-9627 4Museum Solnhofen, Banhofstraße 8, 91807 Solnhofen

 ISBN 978-3-946705-00-0 *Corresponding author; E-mail: [email protected]

Zitteliana 89, 305–329.

Abstract

The locality of Brunn, Oberpfalz, represents the oldest setting within the area usually included within the Solnhofen Archipelago, dating to the Subeumela Subzone of the Late Kimmeridgian. The locality has yielded a rich vertebrate fauna from eight different levels of plattenkalks, dominated by actinopterygian osteichthyans. Apart from rare chondrichthyans, the fish fauna includes macrosemiid gyn- glimodians, caturid and ophiopsid halecomorphs, as well as a new specimen of the enigmatic halecomorph Ainia, and a diverse array of teleosts. Tetrapods are represented by reptiles, including rare marine turtles, at least two new taxa of rhynchocephalians, an atoposaurid and a teleosaurid crocodyliform, and rhamphorhynchid and pterodactyloid pterosaurs. Whereas the fish fauna shows close affinities with other Kimmeridgian faunas, not only from the Solnhofen Archipelago, but also from other European sites, such as Cerin, a high degree of endemism is striking within the reptile fauna. Key words: Solnhofen Archipelago, Kimmeridgian, Actinopterygii, Rhynchocephalia, Pterosauria

Zusammenfassung

Die Lokalität bei Brunn, Oberpfalz, ist in die Subeumela-Subzone des späten Kimmeridgium datiert und repräsentiert somit die älteste Fundstelle des Solnhofen-Archipels. Die Fundstelle hat eine reichhaltige, von Actinopterygiern dominierte Wirbeltierfauna aus acht ver- schiedenen Plattenkalkhorizonten geliefert. Abgesehen von seltenen Chondrichthyern beinhaltet die Fischfauna Gynglimodier der Familie der Macrosemiiden, Halecomorpha der Familien Caturidae und Ophiosidae sowie ein neues Exemplar des seltenen Halecomorphen Ainia und eine diverse Teleosteer-Fauna. Tetrapoden sind durch Reptilien vertreten, darunter seltene marine Schildkröten, mindestens zwei neue Taxa von Rhynchocephalen, ein atoposaurider und ein teleosaurider Crocodyliforme und rhamphorhynchide und pterodactyloide Flugsaurier. Während die Fischfauna große Ähnlichkeit mit anderen Faunen des Kimmeridgiums, nicht nur des Solnhofen-Archipels, son- dern auch anderer europäischer Fundstellen, wie etwa Cerin, weist die Reptilienfauna einen erstaunlich hohen Endemismus auf. Schlüsselwörter: Solnhofen-Archipel, Kimmeridge, Actinopterygii, Rhynchocephalia, Pterosauria

1. Introduction known, even in the technical scientific literature, is that the fauna that is collectively often called the The Upper Jurassic limestones of southern Ger- “Solnhofen fauna” (or “fauna of the Solnhofen limes- many have long been known for their exceptional- tones”) actually includes components from different ly preserved fossils (e.g. Barthel et al. 1990; Welln- depositional settings and also different stratigraphic hofer 2008; Arratia et al. 2015). Despite 200 years horizons (see e.g. Röper et al. 1996; Röper 2005). In- of collecting and research, new species are still deed, fossils come from about a dozen different de- being discovered frequently (e.g. López-Arbarello & pocenters (the so-called “Wannen”; see Viohl 1985; Sferco 2011; Hone et al. 2012; Rauhut et al. 2012a, Schmid et al. 2005) and span c. 3.5 Ma from the b; Schröder et al. 2012; Ebert et al. 2016), and even Late Kimmeridgian to the Early Tithonian (Schwei- several new localities have been explored lately (e.g. gert 2007, 2015). The thin-bedded to laminated lime- Fürsich et al. 2007; Viohl & Zapp 2007; Ebert & Kölbl- stones of the southern Franconian Alb are thought Ebert, 2008; Heyng et al. 2011). What is less well to have been deposited in a number of depressions Zitteliana 89 306 within a sponge-algal-coral reef environment of an lent knowledge of the geological and stratigraphic archipelago at the northern rim of the Tethys Ocean framework of these different localities (see Meyer (Meyer & Schmidt-Kahler 1990; Keupp et al. 2007; 2015; Schweigert 2015; Viohl 2015a, b, c) provides a Viohl 2015a), recently usually called the Solnhofen unique opportunity to study faunal evolution over a Archipelago. Tropical reef environments and archi- geologically short time in a regional setting. Indeed, pelagos accumulate enormous proportions of global the Solnhofen Archipelago represents a unique win- biodiversity because they evolve as a consequence dow into a Mesozoic shallow marine environment, of highly dynamic geological and climatic processes in which we might be able to study the influence of that generate and constantly modify barriers of dis- local environmental differences and evolution over persal. Thus, the Solnhofen Archipelago potentially a short time scale (possibly speciation or species housed a significant proportion of the Late Jurassic replacement) on faunal composition. An absolutely marine global biodiversity. necessary prerequisite for such studies is a detailed Although the Early Tithonian part of the Altmühltal understanding of the taxonomic composition of Formation in the Solnhofen – Eichstätt area (inclu- the faunas of the different localities / depocenters ding the Eichstätt and Solnhofen members; see / stratigraphic levels concerned. Ongoing syste- Niebuhr & Pürner 2014) is by far the most famous matic studies have already shown cases of small unit and has yielded most fossils so far, this main- clades comprising closely related species endemic ly stems from the fact that these rocks have been to the Solnhofen Archipelago, the evolution of which exploited for commercial purposes for centuries, seems to have been driven by palaeogeographical whereas the overlying Mörnsheim Formation and the and chronostratigraphical changes (López-Arbarello underlying Torleite Formation (and equivalents) are & Schröder 2014; Konwert 2016). as or even more fossiliferous, but have not been fully The locality of Brunn (Oberpfalz) is one of the eas- explored so far. The problem of the mixture of fos- ternmost localities within the Solnhofen Archipelago sils from different depocenters and different strati- (Textfig. 1). It was discovered as a fossil Lagerstätte graphic intervals has been known for more than 100 in the early 1990ies and is protected as an important years (Walther 1904; Mayr 1967), but only recently palaeontological site since 1993, following an official have attempts been made to distinguish the different evaluation by one of the authors (M. Röper). It has faunas and see them in their stratigraphic, geogra- been explored for fossils since 1993 by local enthu- phic and ecological framework (e.g. Röper 2005; Vi- siasts (led by Monika Rothgaenger) in collaboration ohl & Zapp 2007; López-Arbarello & Schröder 2014; with Dr. Martin Röper of the Solnhofen Museum, Ebert et al. 2015). Nevertheless, the generally excel- and collected material is housed in the Bayerische

Figure 1: Map of the area between Solnhofen and Regensburg, with the palaeogeography of the Solnhofen Archipelago and the position of the locality Brunn indicated. Modified from Lane & Ebert (2015). 307 Zitteliana 89

Staatssammlung for Paläontologie und Geologie in Bavarian carbonate platform at the northern margin Munich (see Röper & Rothgaenger 1995; Röper et of the Tethys Ocean (Keupp et al. 2007). This plat- al. 1996; Röper 1997, 2005; Heyng et al. 2015). The form was cut off by the Bohemian Massif in the east, locality of Brunn is especially noteworthy for repre- but connected with further carbonate platforms that senting the oldest fauna of the Solnhofen Archipela- extended along the northern rim of the Tethys from go, having been dated in the Subeumela Subzone of eastern Bavaria to southeastern France (see Ber- the Beckeri Ammonite Zone of the Late Kimmerid- nier & Gaillard 2015; Peyer & Khalloufi 2015). To the gian (Röper & Rothgaenger 1997; Schweigert 2007, north, the shallow epicontinental sea that housed 2015). The only older fossiliferous Plattenkalk locality the south Bavarian carbonate platform was bordered in southern Germany, that of Wattendorf (Fürsich et by the Rhenian Massif. In the Late Jurassic, southern al., 2007; Mäuser 2015), is some 100 km further to Germany was situated substantially further south the north and might therefore not represent the same than today, and thus presented a warm, semi-tropi- archipelago as the depocenters between Solnhofen cal setting, in which extensive sponge-microbial and and Regensburg. The locality of Brunn is thus of coral reefs developed (Viohl 2015a). Due to sea-level special interest for our understanding of the origin of changes, the Plattenkalk facies progressed from the faunas of the Solnhofen Archipelago. east to west from the Upper Kimmeridgian to the The limestones of Brunn are especially rich in Tithonian (Viohl 2015a), with laminated limestones plants and invertebrates, but a number of vertebrate being deposited in shallow depressions (“Wannen”) specimens have also been found (Röper et al. 1996; between the reef complexes. The locality of Brunn Heyng et al. 2015). Only few of the fossils of Brunn is situated at the southern rim of one of the smallest have been scientifically analysed yet, and only two of these depressions, the Pfraundorf-Heitzenhofener vertebrates have been described in detail so far. Arra- Basin. The section of the Brunn quarry exposes c. tia & Schultze (2013) described a new species of pa- 8 m of sediments, presenting an intercalation of chycormiform actinopterygian, Orthocormus roeperi, massive limestone layers and eight finely lamina- which is known from a single specimen. The other ted Plattenkalk sections (Röper et al. 1996; Röper vertebrate published is the only complete pterosaur & Rothgaenger 1997; Heyng et al. 2015). Whereas found, which was recently described as a new taxon the massive limestone layers are almost devoid of of rhamphorhynchid, Bellubrunnus rothgaengeri, by macrofossils, fossil remains were otherwise found Hone et al. (2012). An elasmobranch discovered in throughout the section in the laminated limestones. Brunn was furthermore identified as Palaeoscyllium, The latter layers range from 10 to 80 cm in thickness though not described in detail, by Thies & Leidner and are often very finely laminated, with 1 cm of Plat- (2011). Likewise, a few actinopterygian specimens tenkalk including up to 70 distinct limestone laminae from Brunn were referred to previously known taxa: (Heyng et al. 2015). Water depth and/or hostile con- two specimens referred to Ophiopsis (=Furo) muen- ditions apparently increased from the bottom to the steri by Lane & Ebert (2012; see also Lane & Ebert top of the section, as the frequency of autochtho- 2015), and two specimens referred to Macrosemius nous benthic fauna and trace fossils decreases (Rö- fourneti by Ebert et al. (2016). Two rhynchocephalian per et al. 1996; Heyng et al. 2015). Finds of the index specimens were briefly described in a popular sci- ammonite Sutneria subeumela in both the lower and ence article by Rauhut & Röper (2013). However, none upper part of the section constrain the age of the en- of these specimens has been described in detail. tire locality to the Subeumela Subzone of the Upper The purpose of the present paper is to give an Kimmeridgian (Röper & Rothgaenger 1997; Schwei- overview of the vertebrate fauna from Brunn, as far gert 2007; Heyng et al. 2015). The fossil assembla- as it is known so far. Detailed descriptions and analy- ge from Brunn is diverse and especially noteworthy ses of the respective taxa is beyond the scope of this for its high percentage of plants, which account for work, but it will serve as a starting point for compa- up to 25% of the macrofossils found (Heyng et al. risons of the vertebrate fauna from this locality with 2015). Apart from aquatic dasycladalean green al- that of other sites within the Solnhofen Archipelago. gae, the abundance of land plants is noteworthy, as it indicates a significantly higher terrestrial input than in most other Plattenkalk localities of the Solnhofen 2. Geological and palaeontological setting Archipelago. The invertebrate fauna includes most of the clades to be expected in a marine Jurassic As the geological and palaeontological setting of setting, including brachiopods, gastropods, bival- the locality of Brunn has been described in detail by ves, cephalopods, crustaceans and diverse remains Röper et al. (1996), Röper & Rothgaenger (1997), of echinoderms, although the planktonic crinoid Röper (1997) and Heyng et al. (2015), only a brief Saccocoma, which represents one of the most com- overview will be given here. The locality of Brunn is mon fossils found in the earliest Tithonian lithogra- the oldest and one of the easternmost and the most phic limestones of the Solnhofen-Eichstätt area, has northern locality within the area that is usually inclu- not been identified so far (Röper et al. 1996; Röper ded in the “Solnhofen Archipelago” (Textfig. 1). In 1997; Heyng et al. 2015). Vertebrate fossils are found the Late Jurassic, this area was part of the southern throughout the section in basically all Plattenkalk in- Zitteliana 89 308 tercalations, but the preservation and completeness 3.2 Actinopterygii of specimens varies between different layers. Thus, the excellently preserved pterosaur Bellubrunnus More than a hundred actinopterygian specimens was derived from Plattenkalk layer 6 (Hone et al. have been found in the excavations in Brunn only 2012), whereas layer 1 yielded an isolated pterosaur during the past twenty years. The preservation of humerus (Röper 1997). Likewise, whereas fishes the specimens is variable and ranges from excel- from layer 1 are often disarticulated (see Röper et lently preserved, articulated skeletons to isolated al. 1996; Röper 1997), the upper layers have yielded body parts. More than half of the specimens could some perfectly articulated fish specimens. On the be identified to at least the generic level, but some other hand, the almost perfectly articulated rhyn- 40 specimens were too fragmentary and could, at chocephalians were derived from Plattenkalk layer 2 best, only be referred to higher taxonomic catego- (Rauhut & Röper 2013). ries. Identification at the species level was often dif- ficult, also due to the unresolved species taxonomy Institutional abbreviations: JME, Jura-Museum of many fish genera known from the Late Jurassic of Eichstätt, Germany; MHNL, Museum d’Histoire Na- southern Germany, so the following overview of the turelle de Lyon, France; SNSB-BSPG, Staatliche results mainly refers to genera. A more detailed ana- naturwissenschaftliche Sammlungen Bayerns, Ba- lysis of the fish fauna of Brunn is currently under pre- yerische Staatssammlung für Paläontologie und Ge- paration by one of the authors (ALA) and co-workers. ologie, Munich, Germany. Except for 28 species of acipenseriforms (stur- geons and paddlefishes), all other living actinopte- rygian taxa (more than 32,500 species according to 3. The vertebrate fauna the most recent counts by Eschmeyer & Fong [2016]) belong to a single clade called Neopterygii. Two main As it is to be expected in a marine setting, acti- lineages are currently recognized within this clade: nopterygian fishes represent the vast majority of the Teleostei, encompassing most of these species, and vertebrate specimens recovered from Brunn. Only Holostei, which is currently represented by only se- few remains of chondrichthyans have been found ven species of gars (Ginglymodi: Lepisosteiformes) so far. Tetrapods are represented by aquatic turtles, and the bowfin, Amia calva (Halecomorphi: Ami- rhynchocephalians, two crocodylomorph remains, iformes). This tremendous asymmetry in the amount and a few pterosaur specimens. of living representatives of the main neopterygian lineages today does by far not reflect the situation 3.1 Chondrichthyes during the early Mesozoic. During the Triassic-Juras- sic the diversity of holosteans probably equaled or Only few chondrichthyan remains have been even exceeded that of teleosts. Both Teleostei and found so far (Heyng et al. 2015). Among them, one Holostei, including the two main holostean lineages complete specimen (Textfig. 2) from plattenkalk layer Ginglymodi and Halecomorphi, are well represented 7 was identified as Palaeoscyllium ? sp. by Thies & in the plattenkalks of Brunn. Leidner (2011: 78), who also commented briefly on the morphology of its dermal denticles, but has yet to be described in detail.

Figure 2: Elasmobranch Palaeoscyllium sp. from Brunn. Scale bar is in 1 cm increments. 309 Zitteliana 89

3.2.1 Holostei: Ginglymodi were referred to M. fourneti by Ebert et al. (2016) in the most recent revision of this genus. The taxonomy Two main lineages of Ginglymodi are currently of Macrosemius species is mainly based on meris- recognized: Lepisosteiformes and Semionotiformes tic features (e.g., number of dorsal fin rays, number (López-Arbarello 2012; López-Arbarello & Wencker of dichotomizing vertical scale rows between dorsal 2016). Only one lepisosteiform genus, Scheenstia fin and lateral line), but M. fourneti is further charac- López-Arbarello & Sferco, 2011 (including at least six terized by the series of enormous scutes protecting species; López-Arbarello 2012) is represented with the caudal peduncle ventrally, between the anal and two species in the Solnhofen Archipelago (Scham- caudal fins, which are particularly well preserved in haupten, Kelheim, Eichstätt and Langenaltheim), SNSB-BSPG 1993 XVIII VFKO–B 27 (Textfig. 3B). but secure records of lepisosteiforms have not been found in Brunn so far. Röper (1997: 210) mentioned 3.2.2 Holostei: Halecomorphi fragments of Lepidotes from plattenkalk layer 7, but we have not seen this material and the presence of As mentioned above, there is a single living hale- Lepisosteiformes has thus yet to be confirmed. comorph species today, Amia calva (bowfin) which Conversely, semionotiforms are much better rep- is restricted to North American freshwaters. Hale- resented in the Solnhofen Archipelago. With the comorphs however, are broadly distributed in both exception of a few Middle Triassic species of still freshwater and marine environments from the early uncertain affinities, semionotiforms are classified in Early Triassic to Recent. Halecomorphs are classi- three families: Semionotidae, including freshwater fied in two orders: the direct lineage to Amia is the species from the Late Triassic and Early Jurassic of Amiiformes and its sister lineage is the Ionoscopi- Europe, South Africa and North America, and Cal- formes (Grande & Bemis 1998). Both amiiforms and lipurbeckiidae and Macrosemiidae, both including ionoscopiforms are present in Brunn. a very interesting diversity of marine taxa, ranging Although the two amiiform families Amiidae, with from the Middle Triassic to the middle Cretaceous two genera Amiopsis and Solnhofenamia, and Ca- of Europe, Africa and North America. The evolution turidae, with three genera, Caturus, Amblysemius of these two families seems to be directly related to and Liodesmus, are represented in different basins the development of the Tethys and the opening of of the Solnhofen Archipelago, amiids have not yet the Atlantic ocean. Both callipurbeckiids and mac- been recorded in Brunn so far. In contrast, Brunn rosemiids are well represented and diverse in most amiiforms include relatively large and complete ca- of the Solnhofen basins, but only two macrosemiid turid specimens. Among them, SNSB-BSPG 1993 genera have been found in Brunn so far: Notagogus XVIII VFKO–B 30 is very poorly preserved and it has and Macrosemius (Textfig. 3). not yet been possible to identify it at the generic le- One of the macrosemiids found in Brunn is the vel. Two other specimens, SNSB-BSPG 1993 XVIII genus Notagogus, with two complete specimens: a P3 and SNSB-BSPG 1993 XVIII P4, which are com- juvenile SNSB-BSPG 1993 XVIII P1, and a complete- plete and excellently preserved, most probably re- ly ossified, probably adult specimen SNSB-BSPG present a new species of Caturus (Textfig. 4A). The 1993 XVIII P2 (Textfig. 3A). Although a detailed study specimen SNSB-BSPG 1993 XVIII P5 (Textfig. 4B) is necessary to properly identify these fishes, the with a standard length (SL) of c. 45 cm represents a largest specimen agrees in body proportions with new and extraordinary large species of Amblysemius the species N. inimontis according to Bartram (1977). (other species in this genus reach up to c. 30 cm SL; Macrosemius is represented by two complete and Lambers 1994). very well-preserved specimens (SNSB-BSPG 1993 Ionoscopiform specimens from Brunn were re- XVIII VFKO–B 23 and VFKO–B 27; Textfig. 3B), which ferred to the Kimmeridgian species Ophiopsis muen-

Figure 3: Macrosemiid Gynglymodii from Brunn. (A) Notagogus sp., SNSB-BSPG 1993 XVIII P2. (B) Macrosemius fourneti, SNSB-BSPG 1993 VIII VFKO–B 27. Scale bars are in 1 cm increments. Zitteliana 89 310 steri (Textfig. 4C) by Lane & Ebert (2012). This species actinopterygians. The scales of Ainia armata are is more widely known under the generic name Furo also arranged in oblique rows, but these rows are Gistel, 1848 and the generic name Ophiopsis Agas- oriented in posterodorsal to anteroventral direction siz, 1834, is normally associated with other species in the dorsum and in posteroventral to anterodorsal (e.g. O. procera, O. attenuata). The nomenclature direction in the ventrum of the caudal peduncle. This of these taxa was recently revised by Lane & Ebert very peculiar kind of squamation is unique to Ainia (2015), who fixed the name Ophiopsis on the spe- armata among actinopterygians. SNSB-BSPG 1993 cies Ophiopsis muensteri Agassiz, 1834, and created XVIII P6 is by far the most complete and best pre- the new name Ophiopsiella for the species Ophiopsis served specimen of this very rare species, making procera Agassiz, 1843, and other closely related spe- a detailed study of this enigmatic taxon possible cies. To our current knowledge the genus Ophiopsi- (López-Arbarello in prep.). ella includes at least seven European and one African species, ranging from the Middle Triassic to the Early 3.2.3 Teleostei Cretaceous (Lane & Ebert 2015). According to Lane & Ebert (2012) the species Furo orthostomus Agassiz, With 32639 valid living species (Eschmeyer & 1841, Furo normandica Wenz, 1967, and Ophiopsis Fong 2016) Teleostei is the largest vertebrate cla- muensteri Agassiz, 1834 are likely to be congeneric de. Although the monophyly of extant teleosts and under the name Ophiopsis. Several nominal species their close phylogenetic relationships to several of Furo from the Tithonian localities of Bavaria are fossil taxa is generally well established and widely currently under study and might represent additional accepted, the delimitation of Teleostei has been pro- genera (Martin Ebert pers. comm. 2016). blematic (see discussions in De Pinna 1996, Sferco One of the most amazing fossils found in Brunn et al. 2015). Following the long tradition of essenti- is certainly SNSB-BSPG 1993 XVIII P6 (Textfig. 4D). alist thinking in taxonomy (De Queiroz 1994), many Although the bones in the skull are crushed and authors attempted to delimit Teleostei on the basis thus broken, the whole fish is almost perfectly- ar of shared derived traits (e.g. Gosline 1965, Patterson ticulated and extraordinarily well-preserved. The 1973, 1977, Patterson and Rosen 1977, Gardiner et fossil is clearly a specimen of Ainia armata (Wagner, al. 1996, Arratia 1999, 2013). The disadvantages of 1846), which is a rare fish in the Late Jurassic lime- apomorphy-based definitions have been extensively stones of Europe. The species is so far known from discussed (e.g. De Queiroz & Gauthier 1990, De Quei- only several specimens from Cerin (MHNL 15260, roz 1994) and recently De Queiroz (2013) stressed MHNL 15389, MHNL 15469, MHNL 150724, MHNL the feasibility of using a stem-based (branch-based, 20271911, MHNL K203, MHNL K369, MHNL K724, maximum clade) definition to define the name of a MHNL K823), France, and two specimens from Ger- total clade. Apomorphy-based definitions of Teleos- many: the holotype (BSPG AS-I-509) from Kelheim tei depend on the features of the taxa that are added (Moser et al. 2017) and the new specimen from or excluded from the base of the clade [e.g. compare Brunn. Other specimens, such as SNSB-BSPG 1959 Arratia (1999: 323) with Arratia (2013: 115)] and ar- I 427 figured by Grande & Bemis (1998: fig. 422B), bitrarily apply the name Teleostei to one or another represent other species. The species has long been node along the stem to the crown-group. known under the name of Callopterus agassizi Thio- In the same line of thought as De Queiroz (1994), llière, 1858, which was given to the French material De Pinna (1996: 150) proposed a clear and stable (Saint-Seine, 1949, Lambers 1992). However, the ge- stem-based definition of Teleostei as follows: “Tele- neric name Callopterus Thiollière, 1858, is a nomen ostei is here defined … as the largest (i.e. most inclu- nudum (Nomenclator Zoologicus Volume 1: 543) and sive) actinopterygian clade not including either the it was replaced by Ainia Jordan, 1919. On the other Halecomorphi (Amia and close relatives) and/or the hand, Thiollière (1858: 784) and Saint-Seine (1949) Ginglymodi (Lepisosteus and close relatives)”. This listed Lepidotus armatus Wagner, 1846, as a junior total group definition equals the definition applied by synonym of Callopterus agassizi Thiollière, 1858. Patterson (1977) and it is conceptually sound and However, according to the ICZN Wagner’s species stable. De Pinna (1996) further named the clade in- name has priority. Therefore, considering all nomen- cluding all living teleosts and its fossil representa- clatorial rules and changes, the correct name of the tives as Teleocephala, and this node-based definiti- species is Ainia armata (Wagner, 1846). on is stable and independent of our knowledge of a The new specimen from Brunn (SNSB-BSPG species representing the immediate sister taxon of 1993 XVIII P7) shows very nicely the most distinctive the crown clade Teleostei (De Queiroz 2013). feature of this species, which is the very incomple- For the reasons explained above, we accept the te squamation, limited to the dorsum and ventrum node-based clade Teleocephala and the stem-ba- of the caudal peduncle. The scales are very small, sed clade Teleostei of De Pinna (1996). Accordingly, quadrangular to circular and covered with a thick and accepting that they are more closely related to layer of ganoine. These scales are not organized in Teleocephala than they are to holosteans (Gardiner oblique rows oriented in anterodorsal to posteroven- et al. 1996, Nursall 1996, Friedman et al. 2010, Ar- tral direction like the normal body scales of other ratia 2013, Sferco et al. 2015), pycnodontiforms, pa- 311 Zitteliana 89

Figure 4: Halecomorphs from Brunn. (A) Caturus sp., SNSB-BSPG 1993 XVIII P3. (B) Amblysemius sp., SNSB-BSPG 1993 XVIII P5. (C) Ophiopsis muensteri, SNSB-BSPG 1993 XVIII 12. Ainia armata, SNSB-BSPG 1993 XVIII P6 Scale bars are in 1 cm increments. Zitteliana 89 312 chycormiforms, and aspidorhynchiforms are here re- Pachycormiforms are a still controversial and re- garded as teleosts. In this context it is worth noticing latively poorly understood group of fishes that lived that the name Teleosteomorpha proposed by Arratia during the Jurassic and Cretaceous only. They are (2001) to include her Teleostei and all its stem taxa is very abundant in the Early Jurassic and still diverse, redundant because it is equivalent to Teleostei sensu though generally not as well represented in terms of Patterson (1977) or De Pinna (1996). number and state of preservation of the specimens Aspidorhynchiforms and pycnodontiforms are in the Late Jurassic. An exceptionally well-preserved poorly represented in the plattenkalks of Brunn, Late Jurassic pachycormiform is the Brunn speci- although these fishes are frequently found in other men SNSB-BSPG 1993 XVIII-VFKO B16 (Textfig. basins within the Solnhofen Archipelago (Lambers 5B), which is the holotype of the species Orthocor- 1999, López-Arbarello & Schröder 2014). The most mus roeperi Arratia & Schultze, 2013. This complete complete and best preserved aspidorhynchid speci- and almost perfectly articulated specimen preserves men from Brunn (SNSB-BSPG 1993 XVIII P7) most a plethora of anatomical details, which are unknown probably represents a new species of Aspidorhyn- in other pachycormiform taxa and are critical to sol- chus (López-Arbarello & Schröder pers. obs.), which ve the phylogenetic relationships of this very peculiar shows some very interesting features, otherwise only group of fishes. Although O. roeperi is so far only known in Aspidorhynchus arawaki Brito, 1997, from known in Brunn, Arrattia & Schultze (2013) consi- the Upper Jurassic of Cuba. There is furthermore at dered that it more closely resembles O. teyleri Lam- least one specimen of Belonostomus, figured by Rö- bers, 1988, from the Late Kimmeridgian of Cerin, per et al. (1996: Abb. 113). than O. cornutus Weitzel, 1930, from the Tithonian Brunn pycnodontiforms (e.g., SNSB-BSPG 1993 of Langenaltheim, which are the only two other spe- XVIII P8; Textfig. 5A) are identified as Gyrodus he- cies of this genus. Whether this close morphological xagonus (Kriwet 2001). Gyrodus is probably the similarity might reflect close phylogenetic relation- pycondontiform genus with the broadest distribu- ships still needs to be tested in a cladistic analysis. tion, ranging from the Middle Jurassic to the Early Siemensichthys, a rather common teleost in the Cretaceous, and from Europe to the Caribbean and Kimmeridgian and earliest Tithonian, is also relatively the Pacific coast of South America. The species G. abundant in Brunn (Heyng et al. 2015). This genus, so hexagonus is quite common in the Kimmeridgian far endemic to the Solnhofen Archipelago, includes and Tithonian localities of the Solnhofen Archipela- two species, S. macrocephalus Agassiz, 1834, and go, but rare in the Kimmeridgian of Cerin (Lambers S. siemensis Arratia, 2000. Although the specimens 1999, Kriwet 2001). Röper et al. (1996) also mention from Brunn have not yet been studied to the species a specimen of Procinetes, which is a common genus level, the recent studies indicate closest phylogene- found in Cerin (Lambers 1999), but this identification tic relationships between Siemensichthys and two has yet to be confirmed. genera from the Kimmeridgian of Cerin, Ankylopho-

Figure 5: Teleosts from Brunn. (A) Gyrodus hexagonus, SNSB-BSPG 1993 XVIII P8. (B) Orthocormus roeperi, SNSB-BSPG 1993 XVIII VFKO B16. (C) Allothrissops regleyi, SNSB-BSPG 1993 XVIII 42 VFKO C129. (D) Anaethalion cf. cirinensis, SNSB-BSPG 1993 XVIII P9. Scale bars are 10cm (B), 5 cm (C) and 1 cm (D). 313 Zitteliana 89 rus and Lehmanophorus (Arratia 2000, 2013). A rare taxon of basal teleosts is Pleuropholis, which is re- presented in Brunn by at least one specimen, figured by Roeper et al. (1996: Abb. 111). Ichthyodectiforms are particularly well represen- ted in Brunn, including complete and almost fully articulated specimens of at least two species of Al- lothrissops. Based on the diagnosis and description of Nybelin (1964) and the illustration of the holotype in Thiollière (1854), two specimens (SNSB-BSPG 1993 XVIII VFKO C130 and SNSB-BSPG 1993 XVI- II 42 VFKO C129) represent the species Allothriss- ops regleyi (Textfig. 5C), which was previously only known from the Kimmeridgian of Cerin, France. The Brunn specimens, with approximately 230 mm and Figure 6. Partial skeleton of a juvenile marine turtle, SNSB-BSPG 260 mm of standard length are slightly larger than 1993 XVIII VFKO-A1. Scale in mm. the specimens known from Cerin, which reach a standard length of 218 mm (Saint-Seine 1949: 304 died here. Röper et al. (1996) and Röper (1997) men- specimens). Two other specimens (BSPG 1993 XVIII tioned two further, extremely small and obviously lar- VFKO C132 and BSPG 1993 XVIII 42) probably re- gely disarticulated specimens from Plattenkalk layer present Allothrissops mesogaster, which is a well- 1, but these specimens have not been examined for known species from several other Kimmeridgian and this review; furthermore, a partial plastron of an inde- Tithonian localities of the Solnhofen Archipelago and terminate large turtle has been found recently. Both from Nusplingen, but is so far unknown in Cerin. studied specimens are highly incomplete, making Small teleosts are more abundant, although gene- an identification difficult. A very small turtle speci- rally not so complete and well-preserved as the lar- men, SNSB-BSPG 1993 XVIII VFKO-A1 (Textfig. 6), ger fishes listed in the previous paragraphs. Among preserves only the anterior half of the body, the left them, at least four specimens resemble Anaethalion forelimb, the neck, and the lateral rim of the skull. cirinensis Gaudant, 1968, very closely (Textfig. 5D). The carapace of this specimen was only about 50 This species from Cerin is probably closely related mm wide. The poor ossification of the carapace, as to the recently described Ebertichthys ettlingensis well as the unfinished appearance of the limb bones Arratia, 2016, which, together with Ascalabos voithii and the lack of any ossified carpals clearly indicate Munster, 1839, form the family Ascalaboidae Arra- that this specimen represents an early ontogenetic tia, 2016, thus representing an independent teleost stage. Small turtle specimens with separate ribs and clade restricted to the Late Jurassic European car- a poorly ossified carapace from the Upper Jurassic bonate platform along the norther rim of the Tethys. laminated limestones of Germany have traditionally Anaethalion cirinensis and the specimens cf. A. cir- been assigned to Aplax oberndorferi, a species origi- inensis from Brunn differ from E. ettlingensis mainly nally described on the basis of such an early juvenile in the slightly higher number of vertebrae (40 and 43 turtle specimen by Meyer (1843). Although this spe- vs. 38-39, respectively) and the proportionally slight- cies has often been considered to be a synonym of ly shorter caudal vertebral region (40-42% vs. 44%, Eurysternum wagleri (Zittel 1877; Lapparent de Broin respectively). et al. 1996), Anquetin & Joyce (2014) recently did not The most common fish in the Solnhofen Archi- find any evidence for such a referral and noted that pelago, Tharsis dubius, is also the most abundant such poorly ossified turtles should best be regarded actinopterygian in Brunn. However, although most of as indeterminate (see also Joyce 2015). This view is the specimens from Brunn are referable to Tharsis followed here. dubius (Blainville, 1918), many others certainly re- The second specimen, SNSB-BSPG 1993 XVIII present a different and new species of this genus. P10 (Textfig. 7) probably represents a predation vic- More derived teleosts belonging to the crown- tim or the result of scavenging, as more than half of group Teleocephala are also abundant, including nu- the carapace and plastron are missing, although the- merous specimens the salmoniforms Leptolepides re are no clear tooth marks. Most of the left costals and Orthogonikleithrus, at least one specimen repre- and peripherals are preserved, but largely damaged, senting a species of the elopomorph Anaethalion, as is the left side of the plastron, which is largely hid- and a single, but very nicely preserved specimen re- den by the carapace elements, and the left scapula presenting a new genus and species of still uncertain and humerus as well as the neck and skull are also relationships. present. As preserved, the carapace is c. 245 mm long, similar to the size of the type specimen of Eu- 3.4 Testudinata rysternum wagleri (Meyer 1839) and somewhat larger than the specimen of Solnhofia described by Joyce Two partial turtle specimens from Brunn were stu- (2000) and the Eurysternum specimens described by Zitteliana 89 314

Figure 7: Partial skeleton of an indeterminate eurysternid turtle, SNSB-BSPG 1993 XVIII P10. Scale in cm.

Anquetin & Joyce (2014). In contrast to Plesioche- a small fragment of a possible pygal plate preser- lys (Anquetin et al. 2014) and Palaeomedusa (Joyce ved posteromedial to it. This indicates that no pygal 2003), the carapace has well-developed peripheral notch might have been present in this specimen, but fontanelles, and it lacks the strong ornamentation this interpretation should be seen with caution, as that is typical for Platychelys (Cadena & Joyce 2015; the incomplete preservation of the peripheral series Joyce 2015). The skull is short (about 20% of the and the poor preservation of the carapace in gene- length of the carapace), posteriorly broad and seems ral leave some doubt whether the last visible lateral to have a short snout, similar to the situation in Eu- element really represent peripheral 11. Finally, the rysternum (Anquetin & Joyce 2014), but in contrast rather large size of the specimen argues against a to Solnhofia, in which the skull is relatively larger and referral to the rather small Idiochelys, but the possi- has a rather long snout (Joyce 2000). The carapace ble lack of neural plates, which is typical for the latter widens towards the posterior third, as it is also the genus (Joyce 2015), cannot be evaluated, due to the case in Solnhofia (Joyce 2000) and Eurysternum (An- incomplete preservation. quetin & Joyce 2014), but the exact shape cannot In summary, SNSB-BSPG 1993 XVIII P10 seems be established, due to the incomplete preservation. to represent an eurysternid pancryptodire, but can- However, the peripheral fontanelles seem to be rela- not be readily identified in any of the well-known taxa tively smaller than in Eurysternum and the carapace from the Upper Jurassic laminated limestones of as a whole less broad than in Solnhofia. The latter is southern Germany and might thus represent a new also a difference to Tropidemys, which has recently taxon. A more detailed analysis of this specimen and been identified in the Kimmeridgian of Wattendorf probably new finds are necessary to clarify the taxo- (Joyce 2015), but the presence or absence of the di- nomy of the eurysternid from Brunn. agnostic dorsal midline keel of this taxon cannot be evaluated, as the neurals are not preserved. Due to 3.5 Rhynchocephalia the incomplete preservation, most of the diagnostic characters of Eurysternum (Anquetin & Joyce 2014) Four specimens of rhynchocephalians have been also cannot be evaluated. Although the posterior rim recovered from the locality of Brunn so far, and, to- of the last preserved peripheral resembles the ob- gether with the type specimen of the pterosaur Bel- lique margin of the 11th peripheral that forms the rim lubrunnus rothgaengeri, represent the best-preser- of the diagnostic pygal notch in this taxon (Anquetin ved reptile specimens found. All four specimens are & Joyce 2014), close examination of this margin complete and largely articulated, but compressed, reveals that it forms a sutural contact, and there is as it is typical for fossils from the laminated limes-

Figure 8: Specimens of a new rhynchocephalian, similar to Kallimodon. (A, C) complete skeleton (A) and skull (C) of SNSB-BSPG 1993 XVIII 3. (B, D) complete skeleton (B) and skull (D) of SNSB-BSPG 1993 XVIII P11. Scale bars are in cm in A and B, 1 cm in C, and in mm in D. 315 Zitteliana 89 Zitteliana 89 316 tones. Based on a preliminary analysis, at least two lingually. In contrast to most rhynchocephalians, the and possibly three different taxa seem to be repre- tooth row on the palatine seems to be poorly deve- sented by these four specimens, indicating a consi- loped or even absent. The mesial dentary teeth are derable diversity of terrestrial rhynchocephalians in low, with well-developed anterior flanges, but distal this locality. Two of these specimens were described teeth become higher, triangular in outline and with briefly in a popular science article by Rauhut & Röper relatively shorter anterior flanges. The mandibular ar- (2013). ticulation of the mandible shows a small concavity, Two specimens, SNSB-BSPG 1993 XVIII 3 and in contrast to the situation in derived sphenodontids, 1993 XVIII P11 (Textfig. 8), represent some of the lar- which show a longitudinal ridge in this region (e.g. gest rhynchocephalians known from the Late Juras- Rauhut et al. 2012a). sic of southern Germany (see Tischlinger & Rauhut The only other rhynchocephalians from the Upper 2015) and seem to belong to the same taxon. The Jurassic laminated limestones of southern Germany former specimen (Textfig. 8A) has a total length of that reach a similar length as these two specimens c. 380 mm, whereas the latter (Textfig. 8B) is even are the type specimen of Piocormus laticeps (Wag- approximately 460 mm in total length. The limbs ner 1852; Cocude-Michel 1967), a specimen refer- are notably short, with the forelimbs being 57% and red to Sapheosaurus from the collection of Helmut 55% of the length of the presacral vertebral column, Tischlinger (Tischlinger & Rauhut 2015: fig. 840) and and the hindlimbs 92% and 82% of the same length, two undescribed specimens from the Kimmeridgian respectively. The body is elongate and slender. Both of Wattendorf (Mäuser 2015: fig. 1027; Tischlinger specimens are exposed in ventral view, so that only & Rauhut 2015: fig. 838) and Painten (Tischlinger & the ventral side of the skull and mainly the maxil- Rauhut 2015: fig. 839). However, the former three of lary and premaxillary dentition are visible. However, these specimens differ from SNSB-BSPG 1993 XVI- SNSB-BSPG 1993 XVIII 3 has the lower jaw slightly II 3 and 1993 XVIII P11 in the broader, more robust displaced, so that the dentition of the right mandi- body, whereas the specimen from Painten shows a ble is exposed in medial view (Textfig. 8C), and the very different dentition. In general proportions, the left dentary dentition of SNSB-BSPG 1993 XVIII P11 specimens from Brunn are similar to the holotype and is also visible in medial view (Textfig. 8D). The skull a referred specimen of Kallimodon (Table 1; Cocude- seems to have a rather long snout, as it is the case in Michel 1963), but differ from these in the considerab- Kallimodon (Cocude-Michel 1963). As in the Recent ly larger body size (the type of Kallimodon, one of Sphenodon (Robinson 1976) there is only one, large the largest specimens referred to this genus known, premaxillary tooth. No successional teeth (Robinson is c. 180 mm long as preserved, and was certain- 1976; Apesteguía et al. 2012) seem to be present ly no more than 250 mm long when considering the in either maxillary or dentary. The lateral dentition of missing tip of the tail) and the dentition. Thus, these additional teeth is unusual in that it is strongly hete- specimens cannot be referred to any of the known rodont, with a gradual change from small, triangular species of rhynchocephalians from the laminated li- teeth with well-developed posterior flanges anterior- mestones of southern Germany and thus most pro- ly to increasingly transversely expanded, ridge-like bably represent a new taxon, possibly related to Kal- teeth posteriorly, the posteriormost of which resem- limodon. However, as the latter genus is also in need ble the condition seen in opisthodontians in being of revision (Rauhut & López-Arbarello 2016), no new wider transversely than long mesiodistally (see e.g. name is proposed here, pending such a revision. Throckmorton et al. 1981; Martínez et al. 2013; Ape- The specimen SNSB-BSPG 1993 XVIII 4 also re- steguía & Carballido 2014). However, also the most presents a rather large animal, with a total length of distal teeth retain a well-developed posterior flange 345 mm (Textfig. 9). The body is more robust than in

Figure 9: Skeleton of a new sphenodontine rhynchocephalian, SNSB-BSPG 1993 XVIII 4. Scale in cm. 317 Zitteliana 89 the specimens described above, and the limbs re- latively longer, with the hindlimbs slightly exceeding the presacral vertebral column in length; the ratios of limb lengths to presacral vertebral column length are intermediate between that seen in Kallimodon and that of Homoeosaurus (Table 1). As in the specimens described above, the tail is very long, accounting for c. 220 mm of the length of the animal, with the final c. 50 mm being regenerated (see Rauhut & Röper 2013: abb. 9), indicating the capability of tail autoto- my, as it has been described for other Jurassic rhyn- chocephalians (Tischlinger & Wild 2009). As in the specimens described above, the skull is visible in ventral view, with the tooth-bearing ele- ments being slightly disarticulated and the mandi- bles displaced (Textfig. 10). The skull seems to have been relatively shorter and broader than in Kallimo- don and the specimens described above. The den- tition differs markedly from that of other rhynchoce- phalians known from the Late Jurassic limestones of southern Germany (see Cocude-Michel 1963; Figure 10: Skull of a new sphenodontine rhynchocephalian, SNSB-BSPG 1993 XVIII 4. Abbreviations: ld, left dentary; lm, left Carroll & Wild 1994; Rauhut et al. 2012; Rauhut & maxilla; lpal, left palatine; lpm, left premaxilla; rd, right dentary; López-Arbarello 2016). The premaxilla bears three rm, right maxilla; rpal, right palatine; rpm, right premaxilla. Scale teeth, with the mesial two being smaller, incisor-like in cm. and closely appressed, whereas the third is larger, canine-like and separated from the more mesial chocephalians. The anterior end of the dentary also teeth by a deep cleft. The lateral maxillary and den- shows some successional teeth, including two smal- tary dentitions show some remarkable similarities ler mesial teeth and an enlarged, canine-like tooth. to that of the recent Sphenodon (Robinson 1976). Posteriorly follows a row of conical additional teeth, The anterior part of the maxilla bears three to four which gradually increase in size posteriorly, as in successional teeth, with the last of these being en- the modern Sphenodon. The jaw articulation in this larged and fang-like, followed by a small row of very specimen is formed by a longitudinal ridge on the small and largely worn hatchling teeth. Posterior to articular, as in derived sphenodontians. this row, several large, conical additional teeth are This specimen clearly differs from all other rhyn- present. The palate also shows a row of robust, but chocephalians described from the Late Jurassic li- slightly smaller teeth that was apparently parallel mestones of southern Germany and thus certainly to the maxillary tooth row, as in most derived rhyn- represents a new taxon. Its general characteristics,

Table 1: Selected measurements of the rhynchocephalian specimens from Brunn in comparison to other rhynchocephalian specimens from the Late Jurassic of the Solnhofen Archipelago: SNSB-BSPG 1887 VI 502: neotype of Homoeosaurus maximiliani; SNSB-BSPG 1887 VI 1: holotype of Kallimodon pulchellus.

1993 XVIII 3 1993 XVIII P11 1993 XVIII 4 1993 XVIII P12 1887 VI 502 1887 VI 1

Humerus 21,4 22,7 17,4 7,4 18 16 lower arm 14,8 15,9 15,7 5,7 15,5 11 Manus 17,9 21,8 16,1 7,4 16 16 forelimb 57,1 68,6 55,7 23,8 55 46 Femur 29,3 33,2 26,8 9,5 23,5 20,5 lower leg 22,1 22,5 19,6 9,5 22 16,5 pes 32,1 34 27 13,1 27 29 hindlimb 92,9 102,3 81,7 34 78 74 presacral vertebrae 100,7 125,5 73,3 25,5 66 87 Forelimb/hindlimb 0,61 0,67 0,68 0,70 0,71 0,62 forelimb/presacral 0,57 0,55 0,76 0,93 0,83 0,53 hindlimb/presacral 0,92 0,82 1,11 1,33 1,18 0,85 Zitteliana 89 318

Figure 11: Skull (inset) and skeleton of a juvenile rhynchocephalian, SNSB- BSPG 1993 XVIII P12. Scales in mm.

especially of its dentition, resemble the condition in only occurs in the hatchling dentition in Sphenodon Sphenodon and other sphenodontines (e.g. Reyno- (Robinson 1976). Thus, only the last two to three ma- so 1996, 2003), indicating that this specimen might xillary teeth of this specimen might be true additional represent a sphenodontine sphenodontid. However, teeth, also testifying to its early ontogenetic stage. as Apesteguía et al. (2012) pointed out, the dentiti- The dentary teeth are more uniform, being low and on of rhynchocephalians and especially the possible pentagonal in outline mesially and becoming higher, homologies of caniniform successional teeth need more pointed, and triangular towards the back of the further study. Pending a more detailed analysis of tooth row. this specimen, we therefore only refer it to Spheno- As with the other specimens from Brunn, this dontidae incertae sedis here. specimen cannot readily be referred to any of the The smallest specimen, SNSB-BSPG 1993 XVIII known rhynchocephalians known from the Upper P12, is only c. 80 mm long and represents a juveni- Jurassic laminated limestones of southern Germany. le individual, as demonstrated by the generally poor The dentition is clearly juvenile and therefore cannot state of ossification of the limbs, especially the wrist be readily compared to any of the taxa known from (Textfig. 11). The limbs are remarkable long in com- adult specimens. The general proportions, especial- parison to the vertebral column, even more so than ly the remarkable long limbs, resemble the condition in Homoeosaurus, which this specimen most close- in Homoeosaurus. However, as basically nothing is ly resembles in respect to its proportions (Table 1). known about allometric changes during the ontogeny The dentition is highly unusual. The premaxilla bears of Mesozoic rhynchocephalians, a referral to this ta- three to four well separated teeth, and the maxilla xon should be seen as tentative. Given the very young shows a row of conical teeth. Interestingly, for most ontogenetic stage of the specimen, it also cannot be of the length of the maxillary tooth row, the dentition excluded that it might simply represent a juvenile of shows alternating large and small teeth, as it is typi- one of the other taxa present in Brunn, especially that cal for reptiles with active tooth replacement, which represented by SNSB-BSPG 1993 XVIII 4. 319 Zitteliana 89

Figure 12: Remains of an atoposaurid crocodyliform, SNSB-BSPG 1993 XVIII VFKO-A5. (A) slab with several caudal vertebrae, he- mapophyses and osteoderms. (B) detail of a caudal vertebra in lateral view. Abbreviations: he, hemapophysis; os, osteoderm. Scales in mm.

3.6 Crocodylomorpha

One specimen, SNSB-BSPG 1993 XVIII VFKO-A5 represents a crocodyliform (Textfig. 12). The speci- men consists of a small limestone slab containing six mid-caudal vertebrae, some chevrons, and several osteoderms (Textfig. 12A), which are clearly croco- dylomorph. The caudal vertebrae (Textfig. 12B) are very small, with the centrum of the well-preserved anteriormost complete caudal being 8 mm long and 3 mm high. They have low, strongly elongate, am- phicoelous centra that are only slightly constricted in lateral view. The neural arch is low and extends over almost the entire length of the centrum, and the pre- and postzygapophyses overhang the centrum as small, triangular processes anteriorly and poste- riorly. The neural spine is an erect, moderately high, triangular process, in which a slightly thickened cen- tral part is connected to the bases of the zygapophy- ses by well-developed pre- and postspinal laminae. Transverse processes are not present, there is only a small lateral ridge at the basis of the thickened Figure 13: Tooth of the teleosaurid crocodyliform cf. Machimo- central portion of the neural spine. The preserved saurus sp. Scale bar is 5 mm. chevrons are short, V-shaped elements in which the hemal canal is not bridged proximally, but the left terior and posterior laminae (e.g. O’Neill et al. 1981; and right rami remain separated up to their connec- Lio et al. 2012; OR, pers. obs.). The general shape of tion with the vertebrae. The osteoderms are elon- these vertebrae closely corresponds to a caudal ver- gate, with notably convex lateral(?) and straight me- tebra of Alligatorellus described by Schwarz-Wings dial(?) margins. There is a wide, anteriorly rounded et al. (2011), and the shape of the hemapophyses anterior(?) and a smaller, more pointed posterior(?) and the osteoderms, and the size are also consi- process, which are connected by a low, longitudinal stent with an atoposaurid identification (Wellnhofer ridge across the external surface of the osteoderm. 1971; Schwarz-Wings et al. 2011: 199, 201). Thus, The external surface shows the typical groove pat- this specimen is referred to an indeterminate atopo- tern of crocodyliform osteoderms. saurid. The low and elongate shape of the caudal verte- A second, probable crocodyliform specimen, bral centra is typical for neosuchian crocodyliform SNSB-BSPG 1993 XVIII P14, consists of a single, mid-caudal vertebrae, as are the relatively short, strongly worn tooth (Textfig. 13). The crown has a slightly overhanging zygapophyses and the modera- round cross-section and is considerably recurved, tely high, erect neural spine with well-developed an- but the tip is worn down to a flat stump, testifying Zitteliana 89 320 of a durophageous diet. Mesial and distal carinae are absent, but the tooth shows rather well-deve- loped longitudinal striations that are mainly parallel, although some converge towards the apex. As the apex is worn away, it cannot be said whether the striations anastomose apically, as it is the case in Machimosaurus and “Steneosaurus” obtusidens (Young et al. 2014; Foffa et al. 2015). Teeth with a rounded cross-section and well- developed striations are usually found in plesio- saurs (e.g. Massare 1987) and several groups of crocodyliforms, including teleosaurid thalattosu- chians (Vignaud 1997; Mueller-Töwe 2006; Foffa et al. 2015). However, sauropterygian teeth are usual- ly more slender and lack the extensive wear seen in this specimen (Massare 1987). Furthermore, this clade is exceedingly rare in the lithographic limes- tones of southern Germany, with only a single tooth having been reported from the Solnhofen Archipela- go (Wagner 1852). Within crocodyliforms, the mari- Figure 14: Holotype specimen of Bellubrunnus rothgaengeri, ne thalattosuchians are the best represented group SNSB-BSPG 1993 XVIII 2. Scale bar is 1 cm. within the Solnhofen Archipelago (Frey & Tischlinger 2015). Whereas the metriorhynchid genera Cricosau- thus, for which little ontogenetic data exists, whereas rus, Geosaurus and Dakosaurus usually have more O’Sullivan & Martill (2015) discuss that several of the flattened teeth with well-developed, often serrated supposedly diagnostic characters of Bellubrunnus carinae and no or only weakly developed striations rothgaengeri are also found in juvenile specimens of (Young & Andrade 2009; Andrade et al. 2010; Young Rhamphorhynchus, while others are difficult to- ve et al. 2010, 2012), many teleosaurids have well-de- rify and might reflect preservation. The latter authors veloped longitudinal striations on the tooth crowns conclude that „For now, we consider Bellebrunnus (Vignaud 1997). The teeth in most teleosaurids are [sic.] to be sufficiently similar to Rhamphorhynchus more slender and usually do not show a large amount to be considered congeneric based on the current of wear, but the durophageous “Steneosaurus” ob- diagnosis for the genus“ (O’Sullivan & Martill 2015: tusidens and Machimosaurus have stout teeth with 399), but note that two other characters, the lack rounded cross sections and well-developed stria- of stiffening rods formed by the zygapophyses and tions (Young et al. 2014; Foffa et al. 2015). “Stene- the chevrons in the tail and the anteriorly curved last osaurus” obtusidens has weakly developed carinae wing phalanx, indicate that it might represent a spe- in the teeth, while these are missing in at least some cies distinct from Rhamphorhynchus muensteri. The teeth of Machimosaurus, so we tentatively refer this former character, the lack of stiffening rods in the tooth to the latter genus. Although Machimosaurus tail, might also be an ontogenetic character, as the- is a common taxon in the Late Jurassic of Europe se rods seem to be mainly formed by ossified ten- (Young et al. 2014), this is only the second record dons in animals in which such a stiffening of the tail of this genus in the Solnhofen Archipelago, the first is present (e.g. Ostrom 1969). However, the ultimate being a single tooth from the Tithonian Mörnsheim wing phalanx of Bellubrunnus rothgaengeri is indeed Formation of Mühlheim (Heyng et al. 2011). unique in that it is notably curved anteriorly, instead of slightly posteriorly, as it is the case in most pte- 3.7 Pterosauria rosaurs. As these phalanges are preserved in both sides of the wing and show identical morphology, this Given that pterosaurs represent the most com- difference can furthermore also not be accounted for mon group of terrestrial tetrapods in many of the Ba- by preservation, and there is currently no evidence varian Plattenkalk localities, surprisingly few speci- for ontogenetic changes in the curvature of the wing mens have so far been recovered from Brunn. The phalanx in pterosaurs. Thus, we regard the species most remarkably specimen is certainly the complete, B. rothgaengeri as valid, and, given the uncertainty articulated skeleton of a juvenile rhamphorhynchid, of ontogenetic changes in other rhamphorhynchids, SNSB-BSPG 1993 XVIII 2 (Textfig. 14), which was also provisionally retain the genus Bellubrunnus, un- described as a new genus and species, Bellubrun- til ontogenetic studies and/or unambiguously shared nus rothgaengeri, by Hone et al. (2012). Due to the derived characters conclusively show that the spe- early ontogenetic stage of the specimen, its validity cies can be referred to either Rhamphorhynchus or as a distinct taxon has repeatedly been questioned. Scaphognathus. It should be noted, though, that the Thus, Tischlinger & Frey (2015) recently noted that genus Rhamphorhynchus is in need of revision, as the specimen is remarkably similar to Scaphogna- numerous new finds and recent developments in our 321 Zitteliana 89

Figure 15: (A) Anterior end of the snout of a new gnathosaurine ctenochasmatid pterosaur in ventral view. (B) Detail of the dentition, showing marked carina in mesial teeth and enamel ornamentation in more distal teeth. Scale bars are in mm. understanding of pterosaur anatomy, diversity, taxo- tooth bases. The alveoli face laterally and anteriorly, nomy and phylogeny make a re-assessment of the as in Gnathosaurus (Wellnhofer 1970). The preserved taxonomic conclusions of the last major revision of teeth represent mainly or even exclusively maxillary this taxon (Wellnhofer 1975a) necessary. teeth, depending on the position of the premaxillar- Apart from Bellubrunnus, only a few additional maxillary suture. The anterior maxillary dentition is pterosaur specimens have been found, a section of notable for a marked change in tooth morphology conjoined jaws, two isolated appendicular elements from the posteriormost preserved to the first maxilla- and a small assemblage of broken wing phalanges. ry tooth. Although the teeth are generally incomple- Unfortunately, however, the latter assemblage is te, and most lack the tip, the posterior teeth seem poorly preserved, so these bones cannot currently to be lower and are considerably more robust than be identified beyond the level of Pterosauria indet. the anterior teeth. All teeth are pointed and stron- The jaw represents the anterior end of the snout, gly recurved, the anterior teeth with a notable kink including the anterior ends of the maxillae, the pala- at about half-height, basal to which the tooth seems tines, and the premaxillae (Textfig. 15). The snout is to be slightly bulbous, whereas the apical part is very slender, with almost parallel margins in the region slender and pointed. The more distal teeth show a of the anterior maxillae. The tip of the snout expands well-developed, granulate ornamentation on the en- slightly to form a small terminal rosette (Textfig. 15A), amel (Textfig. 15B), similar to the condition found in similar to the situation in Gnathosaurus (Wellnhofer many sauropods (e.g. Holwerda et al. 2015); this or- 1970) and Plataleorhynchus (Howse & Milner 1995), namentation is more weakly developed in the more though less conspicuous. Sutures between the diffe- mesial teeth. The slender mesial teeth have a mar- rent elements are difficult to establish and they might ked mesial carina that is slightly inclined lingually, be partially fused, as in other pterosaurs (e.g. Howse resulting in a longitudinal groove on the lingual side & Milner 1995). Whereas the palatines are separate adjacent to the carina (Textfig. 15B). Judged by the posteriorly, their anterior parts seem to be fused size of the alveoli, the premaxillary teeth seem to and form an anteriorly tapering plate of bone bet- have been somewhat larger than the anterior maxil- ween the maxillae, as in Plataleorhynchus (Howse & lary teeth, though not to the degree seen in Ceara- Milner 1995). Although no clear suture between the dactylus (Unwin 2002). premaxillae and maxillae is visible, the anterior ex- Based on the narrow, parallel-sided rostrum, the tend of the palatines indicates that the premaxillae more laterally than ventrally pointing teeth and the are relatively shorter than in other ctenochasmatids shape of the anterior maxillary teeth, this specimen (Wellnhofer 1970; Dong 1983; Howse & Milner 1995; can be referred to the Ctenochasmatidae (see Welln- Unwin 2002; Bennett 2007a) and probably bore only hofer 1978; Unwin 2002). Within ctenochasmatids, four to five tooth positions, with the anteriormost Unwin (2002) distinguished the Ctenochasmatinae teeth pointing anteriorly. from the Gnathosaurinae (see also Andres & Ji 2008; The right maxilla preserves six to seven tooth po- Andres et al. 2014), with the latter being characte- sitions. The rims of the alveoli are somewhat expan- rized by a rounded expansion of the anterior end of ded, forming a “collar” for the tooth, as in Platale- the rostrum. The fact that the specimen from Brunn orhynchus (Howse & Milner 1995) and, to a lesser shows such an expansion, though less well deve- degree, in Gnathosaurus (Wellnhofer 1970), giving loped than in Gnathosaurus and Plataleorhynchus, the rostrum a slightly festooned appearance. The indicates that it might be referable to that subfamily. teeth are relatively widely spaced, with the distance However, it differs considerably from all other gna- between alveoli being more than the width of the thosaurines in the short premaxilla with few teeth, Zitteliana 89 322 expanded distal articular end. The latter seems to be twisted in respect to the proximal end, so that it faces slightly anterolaterally. In comparison with the humeri of other pterosaurs known from the Upper Jurassic laminated limes- tones of southern Germany, this element is notably large and robust. Indeed, this element is considerab- ly longer than any humerus of non-pterodactyloid pterosaurs from the Upper Jurassic laminated limes- tones of southern Germany measured by Wellnhofer (1975a). The specimen furthermore differs from the common rhamphorhynchids of the laminated lime- stones of southern Germany in the less expanded deltopectoral crest of the humerus and the also less expanded internal tuberosity at the proximal end (Wellnhofer 1975b). The only non-rhamphorhynchid “rhamphorhynchoid” from the Upper Jurassic of southern Germany, Anurognathus, has a less stron- gly expanded deltopectoral crest, but differs from SNSB-BSPG 1993 XVIII VFKO-A3a in the also con- siderably expanded internal tuberosity (Wellnhofer 1975a; Bennett 2007b). In Bellubrunnus, both the deltopectoral crest and the internal tuberosity are less expanded than in other rhamphorhynchids (Hone et al. 2012), but still more so than in the cur- rent specimen. Furthermore, the condition in Bellu- brunnus most probably reflects the early ontogene- tic stage of the specimen, and adults of this taxon Figure 16: Isolated limb bones of pterodactyloid pterosaurs might have been more similar to Rhamphorhynchus from Brunn. (A) Right humerus of an unidentified pterodactyloid, in humeral morphology. SNSB-BSPG 1993 XVIII VFKO-A3 in posterior view. (B) Right In terms of size, the specimen from Brunn falls wi- femur of Ardeadactylus cf. longicollum, SNSB-BSPG 1993 XVIII P13, in posterior view. Scale in mm. thin the range of the largest specimens of pterodac- tyloids listed in Wellnhofer (1970). Apart from a now the rapid change in tooth morphology in the anterior lost specimen referred to “Pterodactylus” grandis maxilla, and the ornamented enamel in more distal from the Mörnsheim Formation (Wellnhofer 1970), teeth and marked mesial carina in mesial teeth and only one described specimen from the Late Juras- thus certainly represents a new taxon. A formal de- scription of the new taxon will be presented else- where. The two appendicular specimens, an isolated right humerus and an isolated right femur, are remar- kable for their large size (Textfig. 16). The right hume- rus SNSB-BSPG 1993 XVIII VFKO-A3 is 100.2 mm long and robust (Textfig. 16A). The proximal end has a saddle-shaped articular surface, as it is typical for pterosaur humeri. Thus, the articular end is concave transversely, but convex anteroposteriorly. A well- developed, elongate, subrectangular internal tube- rosity is present, but the deltopectoral crest, which was found to be of systematic value in pterosaur phylogenies (e.g. Unwin 2003; Andres et al. 2014), is, unfortunately, largely hidden in the matrix. However, judged by the visible parts, it was most probably rat- her moderately developed, and not strongly expan- ded anteriorly, as it is the case in Rhamphorhynchus and other rhamphorhynchids (Wellnhofer 1975a, b, 1978; Padian 2008; Andres et al. 2010). No pneu- matic foramen is visible on the posterior side of the Figure 17: Proximal end of right femur of Ardeadactylus cf. longi- proximal humerus. The shaft is stout and straight, collum, SNSB-BSPG 1993 XVIII P13. Abbreviation: pf, pneumatic and expands distally towards the strongly laterally foramen. Scale in mm. 323 Zitteliana 89 sic laminated limestones of southern Germany has a longer humerus, an isolated wing from the Solnhofen Formation at Schernfeld (JME 1963 1a) that was re- ferred to as “Pterodactylus” sp. by Wellnhofer (1970), which exceeds the specimen from Brunn by 16 mm. The humerus of this specimen is furthermore similar to SNSB-BSPG 1993 XVIII VFKO-A3a in being stout, having a moderately developed internal tuberosity and deltopectoral crest and gradually expanding distally. Thus, we refer this specimen to an indeter- minate pterydactyloid. Based on the unusually large size, it might represent the same taxon as the femur described below. The right femur SNSB-BSPG 1993 XVIII P13 is 87.6 mm long and more slender than the humerus described above (Textfig.16B). It has a well set-off femoral head that is more proximally than medially Figure 18: Reptilia indet., SNSB-BSPG 1993 XVIII 2/2009, two directed, as it is typical for pterosaurs (Wellnhofer dorsal vertebrae in posterior view. Scale in mm. 1978). The shaft is straight and of subequal width throughout its length in anterior view. The external men to Ardeadactylus cf. longicollum. Unfortuna- trochanter (Wellnhofer 1975b; greater trochanter or tely, in the neotype of Ardeadactylus longicollum, trochanter major of other authors, e.g. Bennett 2001; a specimen from the Kimmeridgian of Nusplingen Cordoniú et al. 2006) is well developed and subrec- (Wellnhofer 1970), the proximal ends of both femora tangular in shape, starting at the lateral side of the are covered by pelvic elements (Plieninger 1907). base of the femoral head and extending for some 9 mm distally. In contrast to many pterosaurs, it does 3.8 Reptilia indet. not fade gradually into the femoral shaft distally, but is clearly offset by an oblique step (Textfig. 17). At One specimen, SNSB-BSPG 1993 XVIII 2/2009 about half-height of the external trochanter, a large (Textfig. 18), cannot currently be confidently iden- pneumatic foramen pierces the base of the trochan- tified in any clade. It consists of two small dorsal ter (Textfig. 17), in a very similar position as a large vertebrae that are poorly preserved and exposed in foramen in Pteranodon, which Bennett (2001) iden- posterior view. Interestingly, the vertebral centra are tified as a nutrient foramen. However, nutrient fora- slightly procoelous, as it is found in eusuchian cro- mina in archosaur femora are usually placed more cyliforms and squamates. The neural arch is rather distally and relatively smaller, so that we consider an high, with the distance between the dorsal rim of the interpretation of this opening as a pneumatic fora- centrum and the transverse processes being similar men more likely. The distal end of the femur is ex- to centrum height. Transverse processes are stron- panded, more so medially than laterally. Unfortuna- gly developed and laterally and slightly dorsally di- tely, the articular end is eroded, so nothing can be rected. No details of the zygapophyseal articulations said about the morphology of the distal condyles. are preserved. The neural spine is rather low (lower Whereas the strongly offset, proximally directed than the neural arch) and posterodorsally directed, femoral head characterizes this specimen as a pte- being hook-shaped, pointed dorsally. Although the rosaur element, the rectangular external trochanter is shape of the vertebral centrum, the stout transverse unusual, especially in respect to the angular appea- processes and the relatively low, posteriorly inclined rance of the distal end of the trochanter. In the vast neural spine indicate a terrestrial reptile, little can be majority of pterosaurs, including Rhamphorhynchus said about the affinities of these elements. In respect (Wellnhofer 1975b), Bellubrunnus (Hone et al. 2012), to size, these vertebrae, which are approximately Anurognathus (Bennett 2007b; Tischlinger & Frey 11-12 mm high, would fit an atoposaurid crocody- 2015), Pterodactylus (e.g. SNSB-BSPG 1937 I 18), lomorph (Wellnhofer 1971; Frey & Tischlinger 2015), Germanodactylus (SNSB-BSPG AS I 745, SNSB- but would be rather large for any squamate known BSPG 1892 IV 1), Aurorazhdarcho (Frey et al. 2011), from the laminated limestones of southern Germany and Cycnorhamphus (Bennett 2013a), the external so far (see Tischlinger & Rauhut 2015). trochanter fades gradually into the femoral shaft di- stally. A morphology similar to that seen in the speci- men from Brunn has so far only been illustrated for a 4. Discussion specimen of Ardeadactylus longicollum (Wellnhofer 1970: Abb. 11b; see also Bennett 2013b). Although Despite the rather short time since the discove- the femur of the latter taxon seems to differ in the ry and excavations of the locality Brunn, this site slightly more elongate and less strongly proximally has yielded a remarkably diverse vertebrate fauna, directed femoral head, we therefore refer this speci- reflecting the much higher density of fossils inthe Zitteliana 89 324 Plattenkalk intercalations at this locality than in the Altmühltal Formation, as pointed out by Röper et al. (1996) and Röper (1997). The vast majority of speci- mens recovered yet need detailed study and analy- sis, but we may offer some preliminary thoughts on the significance of the fauna here. The general composition of the fish fauna of Brunn, with a great majority of small teleosts and less represented aspidorhynchiforms, pachycormiforms, pycnodontiforms, ginglymodians, and halecomor- phs does not seem to differ significantly from other localities within the Solnhofen Archipelago (see e.g. Ebert et al. 2015), except for the scarcity of chond- richthyans, which are relatively abundant in other sub-basins. A closer look at the taxonomic compo- sition reveals, however, significant differences with the Tithonian localities, and close resemblance with other Kimmeridgian localities. The only identified shark, Palaeoscyllium, repre- sents a taxon that was originally described from the Solnhofen Member of the Altmühltal Formation (Wagner 1857), but even the type species, P. formo- sum, also has a wide distribution in the Kimmerid- Figure 19: Proximal end of the femur of cf. Ardeadactylus sp. from the Mörnsheim Formation of Daiting, SNSB-BSPG 1977 XIX 39. gian of Europe (Kriwet & Klug 2004). Furthermore, Abbreviation: pf, oneumatic foramen. Scale bar is 1 cm. other species of this genus range from the Batho- nian to the Aptian (Kriwet & Klug 2004), so, pending & Schröder (2014) for the genus Aspidorhynchus. a detailed study and identification of the specimen Concerning the reptile fauna, a rather high level of from Brunn on the species level, nothing can be said endemism of the Brunn fauna is striking. None of the about its evolutionary significance. However, the ra- more complete, articulated reptile specimens can be rity of the fully marine chondrichthyans supports the referred to any species known from other localities interpretation of Brunn as a rather isolated, margi- within the Solnhofen Archipelago. This is the case nal marine, probably lagoonal environment (Röper & of the eurysternid turtle, which probably represents Rothgaenger 1997; Heyng et al. 2015). a new taxon, although further study of this material The association of actinopterygian fishes from is necessary to confirm this. In the rhynchocephali- Brunn seems to be more similar to that of the Kim- ans, at least two new taxa are clearly present, one meridgian limestones of Cerin (Wenz et al. 1994) and represented by the specimens SNSB-BSPG 1993 other Kimmeridgian faunas of the Solnhofen Archi- XVIII 3 and SNSB-BSPG 1993 XVIII P11, and the pelago (e.g. Painten, Kelheim, Ettling) than to the other by SNSB-BSPG 1993 XVIII 4. Whereas the for- „typical“ Solnhofen fauna in the vicinities of Eich- mer taxon seems to be similar and probably related stätt and Solnhofen, as already suggested by Rö- to Kallimodon (see Cocude-Michel 1963; Rauhut & per (1997). Many of the taxa that could be identified Röper 2013), the second specimen does not show on species level are otherwise mainly or exclusively close resemblance to any other rhynchocephalians found in Kimmeridgian localities. This is the case known from the Late Jurassic of Europe, and seems with the halecomorph Ophiopsis muensteri, which is to rather resemble sphenodontines on the lineage also present in Kelheim, Ettling and Cerin (Lane & towards the recent Sphenodon, such as Cynos- Ebert 2012). Likewise, the halecomorph Ainia armata phenodon (Reynoso 1996; see also Cau et al. 2014). is also present in Kelheim and Cerin. Macrosemius The juvenile specimen can also not be referred to fourneti is found in Cerin and the lower, Kimmeridgi- any known taxon, although it cannot be ruled out an section of the locality of Painten, and differs from that this is due to its early ontogenetic stage. As no- Macrosemius rostratus from the Tithonian localities. ted above, it may also be possible that it represents In respect to the teleosts, Allothrissops regleyi is also a juvenile of one of the other two taxa from Brunn, present in Cerin, whereas Allothrissops mesogaster especially SNSB-BSPG 1993 XVIII 4, to which it is (Agassiz, 1834) is the species represented in the Ti- more similar in respect to its general proportions and thonian localities. Likewise Anaethalion cirinensis is the number of premaxillary teeth. However, both of a species that is also known from Cerin and Ettling. A these characters might be subject to ontogenetic similar actinopterygian fauna also seems to be pre- variation (see Robinson 1976). sent in the Kimmeridgian of Nusplingen (ALA, pers. In the pterosaurs, Bellubrunnus rothgaengeri obs.), further indicating that there was some faunal seems to represent an endemic taxon that has so change in actinopterygian faunas from the Kimmeri- far only been identified in Brunn, and the isolated dgian to the Tithonian, as argued by López-Arbarello gnathosaurine rostrum also represents a new taxon 325 Zitteliana 89 that has otherwise not been reported from the Soln- that is currently dominated by actinopterygian fis- hofen Archipelago. The second identified taxon, Ar- hes, most notably teleosts. The aquatic fauna shows deadactylus, is, however, present in other localities. remarkable similarities to other known Kimmeridgian Wellnhofer (1970) referred a total of six specimens localities from the Late Jurassic carbonate platform to Ardeadactylus longicollum (Pterodactylus longi- along the northern rim of the Tethys. Especially no- collum in Wellnhofer 1970), ranging from the Kim- teworthy is that there might be greater similarity of merdigian of Nusplingen to the Tithonian Mörnsheim this fauna to the fish fauna from the Kimmeridgian of Formation of Daiting. However, three of these speci- Cerin than to the Tithonian faunas of the Solnhofen mens were lost during WW II, and the referral of one Archipelago, indicating that stratigraphic differences other specimen, consisting only of a femur and in- in faunas might be more marked than geographic complete tibia, is doubtful (Bennett 2013b). As only and maybe (at least up to a certain level) ecological the original illustrations of the lost specimens exist, differences. Especially the terrestrial reptiles, main- only two specimens can thus currently be regarded ly represented by rhynchocephalians, furthermore as being securely referable to this species (Ben- show a high degree of endemism. This might be due nett 2013b). Interestingly, both of these specimens to island speciation (Losos & Ricklefs 2009), in which come from Kimmeridgian localities, the neotype rhynchocephalian faunas rapidly diversified after dis- from the hoelderi horizon of the Ulmense Ammonite persal over the different islands of the Solnhofen Ar- Sub-zone of Nusplingen and the referred specimen chipelago, as it has often been interpreted in Recent from the rebouletianum horizon of the Ulmense Sub- lizards species (e.g. Glor et al. 2005). However, rapid zone of Schamhaupten (Wellnhofer 1970; Schwei- replacement of lepidosaurian faunas over geological gert 2015). Being derived from the Subeumela Sub- time can also currently not be ruled out. zone of the Late Kimmeridgian, the record from the The exceptional richness of the locality Brunn in locality Brunn is thus the currently oldest record of combination with the excellent stratigraphic control the genus Ardeadactylus (Schweigert 2015). Given of the finds (i.e. that each specimen can be exactly that all specimens referred to this genus so far come placed within the different Plattenkalk layers) provide from the Kimmerdigian begs the question whether an exceptional opportunity to study the dynamics of this might also indicate some temporal stratigraphic the early stages of faunal evolution in the Solnho- segregation of the pterosaur faunas in the Solnho- fen Archipelago. Furthermore, some layers have only fen Archipelago. This might be contradicted by the been recently explored, such as the plattenkalk la- now lost original specimens of Ardeadactylus longi- yer 2, which has yielded the exceptionally preserved collum (Wellnhofer 1970). Furthermore, there is an rhynchocephalian specimens and thus shows great undescribed specimen from the locality of Daiting potential for other terrestrial components of the fau- in the Mörnsheim Formation (Moernsheimensis na, so important new discoveries from this locality Subzone of the Lower Tithonian; Schweigert 2015) are more than likely. Thus, further exploration of the in the collections of the BSPG (SNSB-BSPG 1977 locality of Brunn, as well as in-depth studies of the XIX 39) that consists only of the sacrum, pelvis and remains that have already been retrieved will certain- left hindlimb of a very large pterosaur. This speci- ly result in important new insights into the origin of men, which is currently identified as “Pterodactylus the Late Jurassic vertebrate faunas of southern Ba- grandipelvis”, shows the same, angular external tro- varia and their relation to other Late Jurassic faunas. chanter in the femur (Textfig. 19) that is also seen in the specimen from Schamhaupten (Wellnhofer 1970) and the specimen from Brunn. As the specimen with Acknowledgements a femoral length of c. 100 mm also falls within the size range reported for Ardeadactylus longicollum First and foremost, we have to thank the Bil- (Wellnhofer 1970), it is probably also referable to this dungs- und Dokumentationszentrum Ostbayerische genus. However, as with the femur of the Scham- Erdgeschichte e.V., who helped with the excavati- haupten specimen, there are also slight differences on both financially and by providing volunteers. We to the femur from Brunn, most notably in the position thank the preparators who have prepared the mate- of the pneumatic foramen, which is more proximally rials, especially Lisa Velser, Martin Kapitzke and Pino placed in SNSB-BSPG 1977 XIX 39 (Textfig. 19). If Völkl, furthermore Helmut Tischlinger for providing these differences may indicate species differences photographs of Aurorazhdarcho, Martin Ebert for or fall within the normal variation within Ardeadacty- enlightening discussions about the fish fauna, and lus longicollum will have to be shown in a revision of Walter Joyce for comments on the turtle specimens. this genus. Günter Schweigert provided critical comments on the manuscript, which helped to improve its quality. Last, but not least, we want to dedicate this work 5. Conclusions and outlook to Winfried Werner, whose dedication to the BSPG has helped in so many ways over the years. This The locality of Brunn has yielded a diverse and, at study was supported by the DFG under project RA least in parts, excellently preserved vertebrate fauna 1012/14-1 (to OR). Zitteliana 89 326 6. References Paläontologie, Abhandlungen 245, 23–31. Bennett SC. 2007b. A second specimen of the pterosaur Anurog- Agassiz L. 1833–1845. Recherches sur les poissons fossils, Tome II. nathus ammoni. Paläontologische Zeitschrift 81, 376–398. Recherches sur les poisons fossiles 1–5. Neuchâtel, Petitpierre Bennett SC. 2013a. The morphology and taxonomy of the pterosaur et Soleure, 1420 p. Cycnorhamphus. Neues Jahrbuch für Geologie und Paläonto- Agassiz L. 1834. Abgerissene Bemerkungen über fossile Fische. logie, Abhandlungen 267, 23–41. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Bennett SC. 2013b. 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